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Nutrient distribution in natural habitats is usually patchy in space and time, however most knowledge about plant growth and behaviour is based on experiments conducted under spatially homogenous conditions. Evidence has accumulated that the growth and competitive interaction of plant species are strongly affected by heterogeneous rather than homogeneous resource distribution, even when the total resource supply remains similar. For this study it is hypothesized that infestations of grasslands with the nitrophilous weed Rumex obtusifolius L. (broadleaved dock) are partially the consequence of its ability to exploit spatial nutrient heterogeneity. This was tested in a full-factorial pot experiment with homogeneously or heterogeneously distributed nitrogen and/or potassium at either normal or increased soil moisture where R. obtusifolius was grown together with three other grassland species (grass: Arrhenatherum elatius L., non-leguminous herb: Taraxacum officinale Weber, leguminous herb: Trifolium repens L.). The species differed significantly in their root morphology (root length and diameter, specific root length, number of root tips) and biomass allocation response to nutrient distribution, as well as to the nutrient type used to create patches and to soil moisture (e.g., significant species × treatment interactions). Generally, the root system of A. elatius showed the highest plasticity to imposed treatments, followed by T. officinale, R. obtusifolius and T. repens. Unexpectedly, root morphology of R. obtusifolius was unresponsive to soil heterogeneity and less responsive to nutrient type and irrigation than that of the other species. Nutrient type used to create patches influenced the biomass allocation to the root system of R. obtusifolius while nutrient distribution and irrigation showed no effect on biomass allocation. Exploitation of soil nitrate-nitrogen and potassium was similar among species but exploitation was individually affected by nutrient type, nutrient distribution and irrigation suggesting that species-specific differences in nutrient storage capacities in roots or adjustments regarding root nutrient uptake kinetics may play an important role. Results from this study show that R. obtusifolius does not seem to have superior traits to utilize soil nutrient heterogeneity, certain nutrient types or higher soil moisture that differentiates it from the other grassland species tested. The observed effects might have consequences for the long-term competitive relationships between species in the grassland community suggesting that cultural and biological management measures oriented towards the improvement of the competitive ability of co-occurring grassland species might also be important in heterogeneous soils.
The paper describes the hydrographic-hydrochemical development in the eastern Gotland Basin between the major saltwater inflows into the Baltic Sea in 1993 and 2003. This period is characterised by only low inflow activity. The most important hydrographic events were the effects of the very strong inflow in 1993 and the weak inflows in 1993/1994 and 1997. The 1993/1994 inflows led to deep-water renewal, a steep fall in deep-water temperatures, and increasing salinity. The effects of the inflow of very warm, saline and oxygen-rich water in autumn 1997 were observed in the deep water in 1998, resulting in temperatures rising to 7◦C. The recent renewal in spring 2003 is reflected in the decreasing temperature, higher salinity and improved ventilation of the bottom water. Changes in the redox conditions exert a considerable influence on the nutrient distribution. During stagnation periods, there is enrichment of phosphate and ammonium, while nitrate is absent. Thus, around 31 μmol l−1 ammonium and 7 μmol l−1 phosphate were measured prior to the water renewal in 2003. Deepwater ventilation results in lower phosphate concentrations of around 2 μmol l−1, the nitrification of ammonium and the occurrence of nitrate. For the observation period, an estimate of nutrients stored in the deep water was done for the eastern Gotland Basin. During the recent stagnation period, there was an increase of up to 150% in the phosphate pool below the halocline, whereas the pool of inorganic nitrogen compounds decreased to 80% compared with 1992 when the previous stagnation period had ended. Under specific circumstances, these unbalanced nutrients can be made available to the upper water layers and can induce large-scale blooms of algae, especially of cyanobacteria.
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